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Taxonomy of Rheophilic Luciobarbus Heckel, 1842 (Actinopterygii,

Cyprinidae) from Morocco with the description of two new species

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DOI: 10.3989/graellsia.2016.v72.153

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 Graellsia, 72(1): e039
enero-junio 2016
ISSN-L: 0367-5041
http://dx.doi.org/10.3989/graellsia.2016.v72.153

TAXONOMY OF RHEOPHILIC LUCIOBARBUS HECKEL, 1842

(ACTINOPTERYGII, CYPRINIDAE) FROM MOROCCO
WITH THE DESCRIPTION OF TWO NEW SPECIES

I. Doadrio1*, M. Casal-López1, S. Perea1, & A. Yahyaoui2

1
Biodiversity and Evolutionary Group, Museo Nacional de Ciencias Naturales, CSIC.
C/José Gutiérrez Abascal, 2, 28006 Madrid, Spain
ID: http://lsid:zoobank.org:author:1514FE9E-2AA2-46D1-BB43-51F5E8EF1566
MC-L: http://lsid:zoobank.org:author:84DEBB1B-A34C-412A-B03B-C3492A4E59BE
SP: http://lsid:zoobank.org:author:75C3112F-5BE9-4512-A9DB-DDA2A10497C7
2
Laboratory of Zoology and General Biology, Faculty of Sciences, Mohammed V. University B.P. 1014 Rabat, Morocco
AY: http://lsid:zoobank.org:author:EA344560-E0FA-4D5D-A51C-23771D124D4B
* Corresponding author: doadrio@mncn.csic.es

ABSTRACT

The genus Luciobarbus Heckel, 1843 is characterized by medium to large fish species occurring in large rivers
and downstream zones with slow-moving waters. Remarkably also rheophilic Luciobarbus occur in Morocco,
which are of small size and exhibit distinct morphological traits as well as different habitat requirements. These
rheophilic species have traditionally been assigned to Luciobarbus nasus (Günther, 1874) and L. magniatlantis
(Pellegrin, 1919), although some authors consider L. magniatlantis as a junior synonym of L. nasus. This lack of
consensus on the taxonomy of rheophilic barbs is constrained by limited population studies that do not encom-
pass their entire distribution range. Using molecular, morphometric, and osteological data we studied popula-
tions of rheophilic barbs of three drainage basins in which they are currently present: Tensift, Moulouya and
Oum er Rbia. Our results clearly identified the rheophilic barbs of each basin as different species. The species
Luciobarbus magniatlantis is a well-recognized species endemic to Tensift Basin. In contrast, the investigated
populations from the Moulouya and Oum er Rbia basins could not be assigned to any previously described
species. Consequently, we describe two new Luciobarbus species in the Moulouya and Oum er Rbia basins.
http://lsid:zoobank.org:pub:2494C25A-F4CB-41A1-B6C5-C124D14FD8C4

Key words: North Africa; Luciobarbus; Systematics; mtDNA; morphology.

RESUMEN

Taxonomía de las especies reófilas del género Luciobarbus Heckel, 1842 (Actinopterygii,
Cyprinidae) de Marruecos con la descripción de dos nuevas especies

El género Luciobarbus Heckel, 1843 se caracteriza por presentar especies de medio y gran tamaño que
viven en las zonas bajas de grandes ríos con aguas lentas. Singularmente, en Marruecos también existen espe-
cies reófilas del género Luciobarbus de pequeño tamaño y que presentan diferentes caracteres morfológicos y
distintos requerimientos de hábitat. Estas especies reófilas han sido tradicionalmente asignadas a Luciobarbus
nasus (Günther, 1874) y L. magniatlantis (Pellegrin, 1919) aunque algunos autores consideran a L. magniatlantis
como un sinónimo de L. nasus. Esta falta de consenso en la taxonomía de los barbos reófilos está limitada por los
escasos estudios poblacionales realizados que no abarcan la totalidad del área de distribución de estos barbos.
Nosotros, usando datos moleculares, morfométricos y osteológicos, estudiamos las poblaciones de los barbos
reófilos de tres cuencas hidrográficas, en las cuales están actualmente presentes: Tensift, Moulouya y Oum er
Rbia. Nuestros resultados claramente identifican a los barbos reófilos de cada una de estas cuencas como dife-
rentes especies. La especie L. magniatlantis es una especie bien definida y endémica de la cuenca del Tensift.
En contraste, las poblaciones analizadas de las cuencas del Moulouya y Oum er Rbia no pudieron ser asignadas
a ninguna especie previamente descrita. Consecuentemente, nosotros describimos dos nuevas especies de
Luciobarbus en las cuencas del Moulouya y del Oum er Rbia.
Palabras clave: Norte de África; Luciobarbus; Sistemática; ADN mitocondrial; morfología.
2 I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui

Recibido/Received: 26/10/2015; Aceptado/Accepted: 2/12/2015; Publicado en línea/Published online: 08/02/2016

Cómo citar este artículo/Citation: Doadrio, I., Casal-López, M., Perea, S. & Yahyaoui, A. 2016. Taxonomy of rheophilic
Luciobarbus Heckel, 1842 (Actinopterygii, Cyprinidae) from Morocco with the description of two new species. Graellsia,
72(1): e039. http://dx.doi.org/10.3989/graellsia.2016.v72.153

Copyright: © 2016 SAM y CSIC. Salvo indicación contraria, todos los contenidos de la edición electrónica de Graellsia se
distribuyen bajo licencia de uso y distribución Creative Commons Reconocimiento no Comercial 3.0. España (cc-by-nc).

Introduction placed the rheophilic L. nasus in the genus Luciobarbus

The genus Luciobarbus Heckel, 1843 comprises
more than 30 species of medium to large size widely
distributed along rivers draining into the Persian
Gulf and the Mediterranean, Caspian, and Black
seas (Doadrio, 1994; Kottelat & Freyhof, 2007).
With approximately 20 species, North Africa and
the Iberian Peninsula show the highest diversity in
Luciobarbus populations, with the genus being one
of the main component of the freshwater fish fauna
of these regions (Doadrio, 1994; Machordom et al.,
1995). Phylogenetic studies using morphological
characters, have provided evidence for Luciobarbus
monophyly (Doadrio, 1990), but monophyly has not
been unambiguously retrieved by molecular mark-
ers, due to the position of the genus Capoeta within
Luciobarbus (Levin et al., 2012; Yang et al., 2015).
The genera Capoeta and Luciobarbus comprise lim-
netic species, most of which migrate upstream to
spawning areas, characterized by the presence of
gravel and clear waters (Banarescu, 1999; Doadrio
et al., 2011). Although they can survive in a variety of
habitats, outside of the reproductive period they are
most abundant in downstream areas. Remarkably, in
the freshwaters of Morocco rheophilic Luciobarbus
species can also be found, which have habitat require-
ments and morphological traits that differ from other
Luciobarbus species (Doadrio, 1990).
The rheophilic species of Luciobarbus are sympatric
with limnetic Luciobarbus and Carasobarbus species
in rivers of the Atlas Mountains in Morocco (Doadrio,
1994), but their habitat is restricted to riffle areas, while
other Atlas Mountain Luciobarbus are more wide-
spread, with preference for slower waters (Doadrio
1990). The rheophilic Luciobarbus in Morocco have
been assigned, so far, to two species: Luciobarbus
nasus (Günther, 1874) and Luciobarbus magniatlantis
(Pellegrin, 1919), which are morphologically charac-
terized, with respect to other Luciobarbus species, by
size rarely reaching 20 cm standard length; thick bar-
bels placed at the extreme anterior of the snout; and
males with equal-sized nuptial tubercles uniformly dis-
tributed over the body (Almaça, 1970; Doadrio, 1990).
This difference in morphology placed L. nasus and
L. magniatlantis in an unresolved taxonomic position,
and they could not be assigned to Luciobarbus when
the genus was formally defined on the basis of osteo-
logical traits (Doadrio, 1990). Subsequently, molecular
studies of mitochondrial genes have unambiguously
together with other North African species (Machordom
& Doadrio, 2001a, 2001b).
Within rheophilic Luciobarbus from Morocco,
the taxonomic position of L. magniatlantis remains
unclear. This species has been traditionally considered
a junior synonym of L. nasus (Estève, 1947; Lévêque
& Daget, 1984). However, several authors reported
morphological traits that unequivocally separate the
rheophilic Luciobarbus into two morphological types
and consider L. magniatlantis as a separate species
(Pellegrin, 1939; Almaça, 1970, 1971; Doadrio, 1990).
So far there were no available studies on the popu-
lation structure of the rheophilic barbs of the four
Moroccan basins in which they occur: Moulouya Basin,
on the Mediterranean slope, and Oum er Rbia, Tensift,
and Kasab basins, on the Atlantic slope (Almaça, 1970;
Doadrio, 1994; Borkenhagen & Krupp, 2013). This lack
of population studies compounds confusion about the
type locality of Luciobarbus nasus and has hindered the
taxonomic work. While the type locality, Tensift Basin,
of L. magniatlantis is not in question (Pellegrin, 1919),
the type locality of L. nasus is unclear, because the two
syntype specimens were preserved together with sea
fishes from Mogador (Essaouira) on the seacoast of
Morocco (Günther, 1874). It is probable that the two
syntypes of Barbus nasus came from material collected
in the Kasab River near Mogador by K. v. Fritsch and
J. Rein in 1872 (Borkenhagen & Krupp, 2013).
We review the taxonomy of rheophilic Luciobarbus
from Morocco, investigating morphology and genetic
traits of specimens from Moulouya, Tensift, and Oum
er Rbia basins, where they have been previously
reported (Almaça, 1970). The recognition of two new
distinct taxonomic entities, lead us to the description
of two new species of Luciobarbus for the freshwaters
of Morocco.
Material and Methods
Our study of Luciobarbus populations was based
on 15 specimens (14 males, one female) from Tensift
Basin, 33 specimens (21 males, 12 females) from
Oum er Rbia Basin, and 12 specimens (12 males)
from Moulouya Basin (Fig. 1; Table 1). Material
from Kasab Basin could not be collected despite con-
secutive efforts over the course of several years. The
Kasab River has been intensively modified in recent
years by anthropogenic pressure as a consequence of
the touristic development of Essaouira (Mogador).

Graellsia, 72(1), junio 2016, e039 — ISSN-L: 0367-5041 – http://dx.doi.org/10.3989/graellsia.2016.v72.153

Taxonomy of rheophilic Luciobarbus from Morocco
Fig. 1.— Sampling localities of Luciobarbus populations in
Morocco. Melloul River, Guercif (1, 2); Moulouya R., Ghafoula
(3); Oum er Rbia R., El Borj (4); Chbouka R., El Herri (5);
Douna R., El Khemis (6); Serrou R., El Herri (7); Arba R., Ksiba
(8); Ourika R., (9); Reraia R., Moulay Brahim (10); N’Fiss R.,
Kasbah Goundafi, and Imaounane (11); Kasab R., Essaouira
(Former Mogador) (12).
Fig. 1.— Localidades de las poblaciones de Luciobarbus
de Marruecos muestreadas. Río Melloul, Guercif (1, 2); Río
Moulouya, Ghafoula (3); Río Oum er Rbia, El Borj (4); Río
Chbouka, El Herri (5); Río Douna, El Khemis (6); Río Serrou,
El Herri (7); Río Arba, Ksiba (8); Río Ourika, (9); Río Reraia,
Moulay Brahim (10); Río N’Fiss, Kasbah Goundafi and
Imaounane (11); Río Kasab, Essaouira (antiguo Mogador) (12).
Three cyprinid species were collected in the Kasab
River during the past two decades: Luciobarbus ksibi
(Boulenger, 1905), “Labeobarbus” reinii (Günther,
1874), and Carasobarbus fritschii (Günther, 1874).
However, the endemic “L”. reinii and L. nasus from
the Kasab River have not been found in the past ten
years, and it is likely that the species have disappeared
or at least they are extremely rare. Therefore, our study
of rheophilic Luciobarbus from the Kasab River was
limited to the museum specimens of the two syntypes
of Barbus nasus.
The material studied comprised the following speci-
mens and locations: Tensift Basin: 3 specimens from
the Reraia River, Moulay Brahim (31.2862, -7.9597),
Morocco (voucher numbers: MNCN 280454-280456);
2 specimens from the N’Fiss River, Imaounane (30.9899,
-8.2011), Morocco (voucher numbers: MNCN 208172-
280173); 7 specimens from the Ourika River, Morocco
(voucher numbers: MNHN-IC-1919-0431-0437);
Lectotype and Paralectotypes of Barbus magniatlan-
tis Pellegrin, 1919; 3 specimens from N’Fiss River,
Kasbah Goundafi (30.9899, -8.2011), Morocco
(voucher numbers: MNHN-IC-1919-0438-040). Oum
er Rbia Basin: 1 specimen from the Douna River, El
Khemis (32.7508, -5.5416) Morocco, (voucher number:
MNCN 55094); 2 specimens from the Serrou River, El
Herri (32.8276, -5.6152), Morocco (voucher numbers:
MNCN 208115-280116); 14 specimens from Arba
River, Ksiba (32.5668, -6.0174), Morocco (voucher
numbers: MNCN: 54987-55000); 9 specimens from
the Oum er Rbia River, El Borj (33.0157, -5.6295),
Graellsia, 72(1), junio 2016, e039 — ISSN-L: 0367-5041 – http://dx.doi.org/10.3989/graellsia.2016.v72.153
3
Morocco (voucher numbers: MNCN 208168-208169;
MNCN 71972-71978); 7 specimens from the Chbouka
River, El Herri (32.8595, -5.6213), Morocco (voucher
numbers: MNCN 279704-279707; MNCN 280083;
MNCN 279695-279696). Moulouya Basin: 3 speci-
mens from the Melloulou River, Guercif (34.2180,
-3.3467 and 34.215, -3.3756), Morocco (voucher
numbers: MNCN 286595-286596; MNCN-290831);
9 specimens from the Moulouya River, Ghafoula
(34.145, -3.388), Morocco (voucher numbers: MNCN
290832-290840). Kasab Basin: 2 syntype specimens
of Barbus nasus Günther, 1874 from the Kasab River
preserved at the Natural History Museum (BMNH
1874.1.30.25-26).
All sampling sites (Fig. 1) presented similar river-
ine morphology, with clear water, rapid current and
gravel bottom.
Twenty-four morphometric measurements were
taken with digital callipers (0.01 mm), and ten mer-
istic variables were counted with a stereoscopic
microscope. The following acronyms were used for
morphometric and meristic characters: A, number of
anal fin rays; AFH, anal fin height; AFL, anal fin length;
APL, anal peduncle length; BL1, first barbel length;
BL2, second barbel length; BD, body depth; BLD,
body least depth; C, central caudal fin rays; CFL, cau-
dal fin length; CPL, caudal peduncle length; D, dorsal
fin rays, DFL dorsal fin length; DFH dorsal fin height;
ED, eye diameter; GR, gill rakers (number); HL,
head length; IOW, interorbital width; LL lateral line
scales; P, pectoral fin rays; PFL, pectoral fin length;
PrAD, pre-anal distance; PrDD, pre-dorsal distance;
PrOL, pre-orbital length; PrPD, pre-pectoral distance;
PrVD, pre-ventral distance; PsOL, postorbital length;
PVL, pectoral-ventral length; RSA, scale rows above
lateral line; RSB scale rows below lateral line; SL,
standard length; V, ventral fin rays; VFL, ventral fin
length; VE, Number of vertebrae. The number of ver-
tebrae was obtained by counting on X-ray images of
specimens from all sampled populations. Osteological
characteristics were investigated through computer
tomography (CT) scan and digital dissection using
VGStudio MAX v2.2 (Volume Graphics, http://www.
volumegraphics.com).
After constructing the measurement matrix,
Burnaby’s method was used to correct for size effect.
The Burnaby method removes the effects of a within-
population size-factor from between-group morpho-
metric analyses through an orthogonal projection
procedure (Burnaby, 1966; Röhlf & Bookstein, 1987).
All analyses were conducted with the corrected
matrix. Morphometric and meristic characters were
analysed independently. To assess sexual dimorphism
and to identify the variables that contributed most
to the variation among populations, two principal
component analyses (PCA) were performed using
the covariance matrix for morphometric characters.
Statistical analyses were carried out using PAST
software (Hammer et al., 2001).
4 I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui

Table 1.— Sampling localities for Luciobarbus from Moroccan and GenBank Accession numbers.

Tabla 1.— Localidades de muestreo para Luciobarbus de Marruecos y números de acceso de GenBank.

Number in GenBank
No. Individuals studied Phylogenetic Accession Number
Population assignment/species Locality Morphology/molecular tree Numbers in map
Moulouya population Melloulou R. Guercif/ 3/1 M3 KU257527 1, 2
Moulouya Basin
Moulouya population Moulouya R. Ghafoula/ 9/3 M1, M2, M4 KU257525, 3
Moulouya Basin KU257526,
KU257528
Oum er Rbia population Oum er Rbia R. El Borj/Oum 9/3 R1, R2, R3 AY004744, 4
er Rbia Basin KU257531,
KU257532
Oum er Rbia population Chbouka R. El Herri/Oum er 7/- 5
Rbia Basin
Oum er Rbia population Douna R. El Khemis/Oum er 1/- 6
Rbia Basin
Oum er Rbia population Serrou R. El 2/1 R4 KU257533 7
Herri/Oum er Rbia Basin
Oum er Rbia population Arba R. Ksiba/Oum er 14/- 8
Rbia Basin
Tensift population. Lectotype and Ourika R./Tensift Basin 7/- xxx 9
Paralectotypes of Barbus magniatlantis
Tensift population Reraia R. Moulay Brahim/ 3/2 T2, T3 KU257536, 10
Tensift Basin KU257537
Tensift population. Paralectotypes of N’Fiss R. Kasbah Goudafi/ 3/- xxx 11
Barbus magniatlantis Tensift Basin
Tensift population N’Fiss R. Imaounane/Tensift 2/2 T1, T4 KU257534, 11
Basin KU257535
Kasab population. Syntypes of Barbus Kasab R. Essaouira/Kasab 2/- 12
nasus Basin
L. ksibi Kasab R. /Kasab Basin. -/2 1, 2 KU257523, 12
KU257524
L. ksibi Reraia R./Tensift basin -/2 3, 4 KU257538, 10
KU257539
L. ksibi Chbouka R./Oum er Rbia -/2 5, 6 KU257529, 5
basin KU257530
L. bocagei Duratón R./Duero Basin -/1
L. comizo Tajo R./Tajo Basin -/1
L. graellsii Irati R./Ebro Basin -/1
L. guiraonis Turia R./Turia Basin -/1
L. microcephalus Zujar R./Guadiana Basin /1
L. sclateri Segura R./Segura Basin /1
L. setivimensis Soumman R./Soumman Basin /1

For molecular analyses, we obtained specimens of
rheophilic Luciobarbus from Oum er Rbia, Tensift,
and Moulouya basins along with the limnophilic
L. ksibi from Oum er Rbia, Tensift and Kasab basins,
(Table 1). Also, the Iberian species of Luciobarbus
and L. setivimensis (Valenciennes, 1842) from Algeria
were added. Aulopyge hueguelli Heckel, 1843 and
Barbus meridionalis Risso, 1827 were selected as
outgroups, based on previous phylogenetic analy-
ses (Zardoya & Doadrio, 1999). Total genomic DNA
was extracted from fin-clip tissue using the commer-
cial kit Biosprint15 for tissue and blood (Qiagen).
For each specimen, the complete region (1140 bp)
of the mitochondrial cytochrome b (cytb) was ampli-
fied. Primers and protocols used for PCR for cytb
followed Machordom & Doadrio (2001b). After check-
ing PCR products on 1% agarose gels, they were purified
by ExoSAP-IT™ (USB) and directly sequenced on
MACROGEN service using a 3730XL DNA sequencer.
Sequences were deposited in the GenBank database
under accession numbers KU257523-KU257539.
Phylogenetic analyses were performed using Bayes-
ian inference (BI) implemented in MrBayes v. 3.2
(Ronquist et al., 2012). The Akaike Information Criterion

Graellsia, 72(1), junio 2016, e039 — ISSN-L: 0367-5041 – http://dx.doi.org/10.3989/graellsia.2016.v72.153

Taxonomy of rheophilic Luciobarbus from Morocco 5

(Akaike, 1973) implemented in jModeltest (Posada,
2008) was used to determine the evolutionary model
that best fit the data. In this case, TIM1+G model was
selected. Bayesian inference was performed using two
independent runs of four Markov Montecarlo coupled
chains (MCMC) of 5×106 generations each to estimate
the posterior probability distribution. Topologies were
sampled every 100 generations, and a majority-rule
consensus tree was estimated after discarding the first
10% of generations. The robustness of the clades was
assessed using Bayesian posterior probabilities. The
average uncorrected p-distances among Luciobarbus
populations were calculated for the cytb gene using
MEGA package v. 6.0 (Tamura et al., 2013).
The number of scales along the lateral line
(median=45), superior transverse line (median=7.5)
and inferior transverse line (median=5.5) was signifi-
cantly lower in the Moulouya population than in the
Tensift and Oum er Rbia (Appendix 1). Scale num-
ber along the lateral line (median=51) and superior
transverse line (median=9.5) were significantly higher
in the Oum er Rbia population than in Tensift and
Moulouya populations.

Results and Discussion

COMPARISON OF MORPHOLOGY AMONG

POPULATIONS

Due to the sexual dimorphism of Luciobarbus

(Doadrio, 1990), and because few female speci-
mens were available, we removed females from the
subsequent morphological analyses. Non-parametric
Kruskal–Wallis and Mann–Whitney post hoc compari-
son analyses were used to assess differences in size
and body shape among populations (Appendix 1). No
differences in SL were found among Luciobarbus pop-
ulations. However we found significant differences in
morphometric measures, with the exception of preven-
Fig. 2.— Morphology of the analysed populations. Arrows
tral and preanal distances, anal and dorsal fin lengths, show principal traits of each population. A: Tensift Basin
pre and post-orbital distances, and caudal peduncle population; B: Oum er Rbia Basin; C: Moulouya Basin. Scale:
length (Appendix 1). 10 mm.
The population from Moulouya basin exhibited an Fig. 2.— Morfología de las poblaciones analizadas. Las
elongated body shape with the smallest body depth flechas muestran los principales rasgos de cada población.
of any studied population. The dorsal fin was placed A: población del Tensift; B: población del Oum er Rbia;
slightly more anterior, and the predorsal distance was C: población del Moulouya. Escala: 10 mm.
shorter, than in Tensift and Oum er Rbia populations.
All fins in the Moulouya population were larger than
the ones present in the remaining populations, with the
height of the anal fin significantly bigger than in other
populations, in some specimens reaching, or nearly
reaching, the anterior rays of the caudal fin when
folded. The population of Oum er Rbia Basin possessed
the longest anal caudal peduncle. This population had a
narrow skull, characterized by the lowest inter-orbital
width. The population of Tensift Basin had the smallest
head, with the head and prepectoral distances shorter
than in Moulouya and Oum er Rbia populations. The
deepest caudal peduncle was also found in the Tensift
population. The fins in the Tensift population were sig-
nificantly smaller than in Moulouya and Oum er Rbia
populations. Some specimens of the Tensift popula- Fig. 3.— Syntypes of Barbus nasus showing the characteristic
tion, belonging to paralectoypes of Barbus magniat- deep caudal peduncle. Figure from the Natural History Museum
lantis, presented three pairs of barbels, a characteristic (London).
that was not observed in any other populations. An Fig. 3.— Sintipos de Barbus nasus mostrando su característico
overview of the general morphology of the analysed y profundo pedúnculo caudal. Figura del Museo de Historia
populations is illustrated in Figs. 2 and 3. Natural de Londres.

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6 I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui

Fig. 4.— Variables that most contributed to the PCA analysis. Dots: Tensift population. Squares: Oum er Rbia population. Triangle:
Moulouya population. Acronyms are defined in Materials and Methods.
Fig. 4.— Variables que más contribuyeron al análisis de PCA. Puntos: población del Tensift. Cuadrados: población del Oum er
Rbia. Triángulo: población del Moulouya. Los acrónimos están definidos en la sección de Material y Métodos.

The principal component analysis (PCA) divided
the specimens into three groups, corresponding to the
populations of the three investigated basins (Fig. 4).
The eigenvalues of the three first principal compo-
nents, with the Burnaby-corrected matrix, explained
most of the variance (Table 2). The highest eigenvec-
tor values (anal fin height and inter-orbital width)
were in agreement with results of Kruskal–Wallis and
Mann–Whitney analyses (Table 2).
OSTEOLOGICAL FEATURES (APPENDIX 2)
As shown by morphometric analyses, the skull of
the Oum er Rbia population was narrower than the
one present in Tensift and Moulouya populations.
In particular, the ethmoid bone was longer and nar-
rower (Appendix 2-1). In contrast, the kinethmoid
bone of the Oum er Rbia population was shorter and
more robust than in other populations (Appendix 2-2).
The posterior branch of the lachrymal bone was elon-
gated in the Oum er Rbia population and shorter in
the Moulouya population (Appendix 2-3). The dentary
bone was shorter in the Tensift population than in other
populations (Appendix 2-4). The anterior process of
the maxilla of the Tensift population was lesser devel-
oped than in Oum Er Rbia and Moulouya populations
(Appendix 2-5). The number of pharyngeal teeth in
Tensift, and Oum er Rbia populations was usually
4.3.2. As in other Luciobarbus species, the juveniles
possessed five teeth in the external row. In the Tensift
population, the fifth tooth was occasionally retained
in the adults. In the Moulouya population, we found
pharyngeal teeth in a 4.2.1 or 4.3.1 configuration
(Appendix 2-6). The Oum er Rbia population had a
thick inner branch of the pharyngeal bone, with reduced
pharyngeal lamina. In the Tensift population, the supe-
rior branch of the pharyngeal bone was strongly flexed
(Appendix 2-7). In Oum er Rbia populations, the last
single ray of the dorsal fin was strongly ossified with
a maximum width of 12-14.5% (median=13.8) of its
height (Fig. 5). In Tensift (6.7-9.5%, median=8.3) and
Moulouya (6.2-9.2%, median=7.4) populations, the
last single ray of the dorsal fin showed a lower level of
ossification (Fig. 5). The Tensift and Moulouya popu-
lations showed weaker denticulations in the last single
ray of the dorsal fin, but higher denticulate density
(Moulouya 3.1-2.4 teeth/mm, median=2.6 teeth/mm;
Tensift 2.2-1.8 teeth/mm, median=2.1 teeth/mm)
(Fig. 5). The population from Oum er Rbia had scarce
and strong denticulations on the last single ray
of the dorsal fin at a density of 1.1-1.4 teeth/mm,
median=1.2 teeth/mm (Fig. 5).
MOLECULAR DATA
The phylogenetic analysis using BI, with Aulopyge
huegelli and Barbus meridionalis as outgroup taxa,
revealed two main clades corresponding primarily to
the Iberian and African species (Fig. 6). The Iberian
group clustered with Luciobarbus setivimensis of
Algeria, as was previously reported (Machordom &
Doadrio, 2001b). Unexpectedly, the populations of
rheophilic Luciobarbus were not monophyletic, and
the Moulouya Basin population was clustered with

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Taxonomy of rheophilic Luciobarbus from Morocco 7

Table 2.— Eigenvalues and eigenvectors for the first three

principal components (PC1-PC3) of 21 morphometric
variables for all rheophilic Luciobarbus populations. Acronyms
are defined in the Material and Methods section. Variables
with the highest eigenvalues for each PC are in bold.

Tabla 2.— Eigenvalores y eigenvectors para los tres primeros

componentes principales (CP1-CP3) de 21 variables
morfométricas para todas las poblaciones de barbos reófilos.
Los acrónimos están definidos en la sección de Material y
Métodos. Las variables con los eigenvalores más altos para
cada CP están en negrita.

Variables PCI PCII PCIII

Eigenvalue 0.0050 0.0019 0.0009
% variance 46.48 18.05 8.85
Eigenvectors
SL −0.2605 −0.0678 −0.1058
Fig. 5.— Denticulation of the last single ray of the dorsal fin in
PrDD −0.0926 0.0432 −0.1735
specimens of the populations from Tensift (A, SL 135.8 mm),
PrPD 0.1747 0.1477 0.0542 Moulouya (B, SL 139 mm), Oum er Rbia (C, SL 126.5 mm)
PrVD −0.0406 0.0515 0.0110 and syntype of Barbus nasus (D, SL 108.7 mm).
PrAD −0.0046 −0.0412 0.0021 Fig. 5.— Último radio sencillo denticulado de la aleta dorsal en
PVL −0.2190 −0.3034 −0.3745 individuos de las poblaciones del Tensift (A, SL 135,8 mm),
Moulouya (B, SL 139 mm), Oum er Rbia (C, SL 126,5 mm)
CPL −0.0301 −0.1112 −0.3078 and sintipo de Barbus nasus (D, SL 108,7 mm).
APL 0.0062 0.1765 −0.3697
BD −0.3888 0.302 0.1404
than the ones reported to sister species of cyprinid
BLD −0.3187 −0.1862 0.0311
fishes (Doadrio & Carmona, 2003, 2006; Doadrio &
HL 0.1575 0.1267 −0.1107 Madeira, 2004; Robalo et al., 2005; Doadrio & Elvira,
PrOL −0.0114 0.0839 0.4119 2007; Domínguez-Domínguez et al., 2007, 2009). In
ED −0.2916 0.2065 0.1496 contrast, genetic distances between Tensift and Oum
PsOL 0.1731 0.1844 −0.0512 er Rbia population were lower than the ones found
PFL 0.2832 0.1257 0.1731 in sister species of Luciobarbus (Casal-López et al.,
VFL 0.2832 0.0867 0.2835 2015).
DFL −0.0053 0.1496 0.1400
DFH 0.2053 0.2113 −0.0097 TAXONOMIC REMARKS
AFL −0.0504 −0.0029 −01324
The three studied populations from the Oum er
AFH 0.5536 −0.3194 −0.1403
Rbia, Moulouya, and Tensift basins possessed molec-
IOW −0.0760 −0.6432 0.4349 ular, osteological, and morphometric traits that iden-
tified them as different species. The assignment of
Oum er Rbia, Moulouya, and Tensift populations to
the Iberian group, while the populations from Oum er the rheophilic species L. nasus was not possible, as the
Rbia and Tensift basins constituted a monophyletic combination of diagnostic traits of L. nasus was not
group with Luciobarbus ksibi from the same basins shared by the studied populations. The combination
and from its type locality (Kasab River). The phyloge- of the diagnostic traits of all populations and the syn-
netical position of the Moulouya population within the types of Barbus nasus are showed in Table 4. The pop-
Iberian group was not resolved. ulation morphologically most similar to L. nasus was
Genetic distances within Moulouya, Oum er Rbia that of the Oum er Rbia Basin. Differences between
and Tensift populations of rheophilic barbels ranged the Oum er Rbia population and L. nasus were found,
from 0.1 to 0.3%. The Moulouya population showed primarily in the morphology of the last single ray of
genetic distances from Tensift and Oum er Rbia pop- the dorsal fin, number of scales in the lateral line,
ulations ranging from 8.1 to 8.4%. The genetic dis- morphology of the pharyngeal teeth, kinethmoid bone
tances between Oum er Rbia and Tensift populations shape and caudal peduncle depth. The maximum
was 1.1% (Table 3). Genetic distances between limno- width of the last single dorsal fin ray of the syntype
philic L. ksibi and rheophilic barbs of the same basins of Barbus nasus was 11.5% of its height, and does not
(Oum er Rbia and Tensift) were from 5.6 to 5.9%. The reach the robustness of that of the Oum er Rbia popu-
genetic distances of Moulouya population with respect lation (12-14.5%, median=13.8) (Fig. 5). The number
to other Luciobarbus species were similar or higher of scales in the lateral line was 46-44 in Barbus nasus

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8 I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui

Fig. 6.— Phylogenetic tree rendered by Bayesian Inference of the mitochondrial cytochrome b gene. Numbers on branches
indicate posterior probability values. Identification of localities is defined in Table 1. Red branches show limnophilic North African
species. Blue branches represent North African rheophilic populations.
Fig. 6.— Árbol filogenético del gen mitochondrial citocromo b obtenido a partir de Inferencia Bayesiana. Los números sobre
las ramas indican valores de probabilidad posterior. La identificación de las localidades está definida en la Tabla 1. Las ramas
marcadas en rojo representan las especies limnófilas del Norte de África. Las ramas marcadas en azul señalan las poblaciones
reófilas del Norte de África.

syntypes and 49-53 scales ( x =51, median=51) in
Oum er Rbia population. Pharyngeal teeth and kineth-
moid bone were more robusts in Oum er Rbia than in
L. nasus (Appendix 2-2, 2-7). Anal peduncle length
was less of two times the BLD while in Oum er Rbia
population was always more of two times.
In the absence of molecular data and a greater num-
ber of specimens for morphological studies, L. nasus
must be considered an endemic species of the Kasab
Basin and L. magniatlantis an endemic species of the
Tensift Basin.
The populations of Moulouya and Oum er Rbia
basins exhibit multiple diagnostic traits differing from
those of L. nasus and L. magniatlantis (Table 4).
DESCRIPTION OF LUCIOBARBUS POPULATIONS

Table 3.— Genetic distances for the complete mitochondrial

The high degree of morphological and genetic dif-
cytb gene. Uncorrected-p genetic distances between species ferentiation of Luciobarbus populations endemic to
are below the diagonal. Uncorrected-p genetic distances the Moulouya and Oum er Rbia basins justifies the
within species are shown in the diagonal. consideration of these population as distinct species.
No available names for these populations exist, and
Tabla 3.— Distancias genéticas para el gen mitochondrial
citocromo b completo. Las distancias genéticas no corregidas
therefore these are described as new species in the
entre especies están debajo de la diagonal. Las distancias present study.
genéticas no corregidas dentro de las especies se muestran
en la diagonal. Luciobarbus guercifensis sp. nov. Doadrio, Perea &
Yahyaoui
Moulouya Oum er Rbia Tensift L. ksibi http://lsid:zoobank.org:act:6DF6332F-F73B-437F-88E1-
Moulouya 0.2 2F4AC7F15F4F
Oum er Rbia 8.4 0.3 HOLOTYPE: Fig. 7, Table 5. MNCN 290835 male, 139.4 mm (SL);
Tensift 8.1 1.1 0.1 Moulouya River, Moulouya Basin, Ghafoula, Mediterranean slope
in Morocco (34.14534, -3.38847) (Fig. 1); 22/6/2015. 378 ATSL,
L. ksibi 9.1 5.9 5.7 0.5
Collected by (Coll.) Doadrio, I; Yahyaoui, A and Perea, S.

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Taxonomy of rheophilic Luciobarbus from Morocco 9

Table 4.— Diagnostic morphological characters of the four populations studied.

Tabla 4.— Caracteres morfológicos diagnósticos de las cuatro poblaciones estudiadas.

Moulouya Oum er Rbia Tensift Kasab

population population L. magniatlantis L. nasus
Width of the last single ray of <10% Height >10% Height <10% Height >10% Height
the dorsal fin
Denticulations of the last single Profusely serrated Sparsely serrated Profusely serrated Sparsely serrated
ray of the dorsal fin 24-26 denticulations 11-13 denticulations 18-22 denticulations 20-21<25 denticulations
in adults in adults in adults in adults
No. scales in lateral line 45-48≤48 49-53>48 51-47≤48 44-46≤48
No. scales above lateral line 6.5-8.5 8.5-9.5 8.5 9.5
Inner row of pharyngeal teeth 1 2 2 2
Pharyngeal Teeth Robust Robust Robust Weak
Lachrymal manubrium Wide Narrow Narrow Narrow
Ethmoid bone Wide Narrow Wide Narrow
Kinethmoid bone Narrow Wide Narrow Narrow
Caudal peduncle Low and elongated. Low and elongated. Short and high. Short and high.
ACL is ≥2.1 the BLD ACL is ≥2.1 the BLD ACL is <2 the BLD ACL is <2 the BLD

PARATYPES: Table 5. MNCN 286595-96: 2 specimens from
Melloulou River, Moulouya Basin, Guercif, Mediterranean slope
in Morocco (34.218035, -3.346732); 9/4/2007. Coll. Doadrio,
I; Doadrio, I jr. and Perea, S. MNCN 290831: 1 specimen from
Melloulou River, Moulouya Basin, Guercif, Mediterranean slope
in Morocco (34.21526, -3.375668). 2/5/2015. Coll. Doadrio, I;
Garzón, P; Perea, S and Yahyaoui, A. MNCN 290832: 1 specimen
from Moulouya River, Moulouya Basin, Ghafoula, Mediterranean
slope in Morocco (34.14534, -3.38847); 2/5/2015. Coll. Doadrio, I;
Perea, S; Garzón, P and Yahyaoui, A. MNCN 290833-290834,
290836-290840: 7 specimens from Moulouya River, Moulouya
Basin, Ghafoula, Mediterranean slope in Morocco (34.14534,
-3.38847); 22/6/2015. Coll. Doadrio, I; Perea, S and Yahyaoui, A.
The holotype and a series of paratypes (12 specimens) have
been deposited at the Fish Collection of the Museo Nacional de
Ciencias Naturales, Madrid, Spain.
DIAGNOSIS: Differs from other known Luciobarbus
species by the following combination of characters:
45-48 scales along the lateral line ( x =45.6, median=45),
6.5-8.5 ( x =7.4, median=7.5) above lateral line, and
5.5 below lateral line. The last single ray of the dor-
sal fin is strongly ossified with the maximum width
6.2-9.2% (median=7.4) of its length. The last single
dorsal fin ray is densely denticulated along its length,
Fig. 7.— Holotype of Luciobarbus guercifensis from the Moulouya
River, Ghafoula, Morocco. MNCN 290835. SL=139.4 mm.
Fig. 7.— Holotipo de Luciobarbus guercifensis del Río Moulouya,
Ghafoula, Marruecos. MNCN 290835. SL=139,4 mm.
and, in adults specimens, the number of denticulations
exceeds 25 (Fig. 6). The ethmoid bone is wider than
its length. Most specimens possess a single pharyn-
geal tooth in the inner row. The number of vertebrae
is 39-41 ( x =39.7, n=11). The interorbital distance is
1.7 to 2.2 times (median=2) the eye diameter. The cau-
dal peduncle is longer than the one seen in L. nasus
and L. magniatlantis, with depth 2.1-2.6 times the
length of the anal peduncle. The posterior segment
(manubrium) of the lachrymal bone is short and high.
Differences in diagnostic characters among analysed
Luciobarbus populations are presented in Table 4.
DESCRIPTION: D III-V 8, A III 5, P I 15-16, V I 6,
C 18; LL 45-48 ( x=45.6, median=45), RSA 6.5-8.5
( x=7.4, median=7.5), RSB 5.5. Pharyngeal teeth in
adults 4.3.1 or 4.2.1. GR 13-15 ( x=14.1, median=14),
VE 39-41 ( x =39.7, median=40). Rarely reaches
200 mm SL. The body is elongated, relative to maxi-
mum body depth compared to other Luciobarbus spe-
cies. Maximum body depth is 16-19% of SL. Head
length ( x =28.5 mm) is 25-27% of SL and greater than
body depth ( x =20.1 mm). The skull is wide, with
the ethmoid bone width greater than its length. The
interorbital distance is 1.7 to 2.2 times eye diameter.
Infraorbital bones are narrow and the lachrymal bone
is shorter than in Oum er Rbia population as conse-
quence of a shorter manubrium. The height of the
manubrium is 20-23% the lachrymal length. Thick lips
and barbels display granular appearance. The first pair
of barbels is short and located at the anterior extreme of
the mouth, reaching the insertion of the second pair of
barbels. The second pair of barbels is thick and reaches
the posterior edge of the eye. The anterior barbel is
23.2-29.7%, and the second 34.6-38.6%, of HL. The
snout is prominent, with preorbital length 10-12.6%
of SL. The preorbital length ( x=13.6 mm) is larger

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10 I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui

Table 5.— Morphometric and meristic measurement of length is 16.6-27.3% of SL. Morphometric and mer-
the holotype and paratypes of Luciobarbus guercifensis. istic measurements for the holotype and paratypes of
Acronyms are defined in the Material and Methods section.
Luciobarbus guercifensis are presented in Table 5.
Tabla 5.— Medidas morfométricas y merísticas del holotipo The colouration of L. guercifensis is silver-yellowish
y paratipos de Luciobarbus guercifensis. Los acrónimos with a paler ventral area (Fig. 2).
están definidos en la sección de Material y Métodos.
DISTRIBUTION: This species is endemic to Moulouya
basin, inhabiting Moulouya and Melloul rivers in riffle
Holotype
MNCN 290835 Paratypes n=11
areas near to the Guercif village (Fig. 1).
Morphometric Measurements Standard
variable (mm) Range Mean Deviation
ETYMOLOGY: The species name ‘guercifensis’ was
selected, as because the species is mainly distributed
SL 139.4 75.7-149.4 109.4 23.5
around the Guercif village in Morocco.
PrDD 66.7 37.1-70.3 52.5 10.4
PrPD 35.2 20.1-38.2 28.3 5.7 COMMON NAME: We propose the English common
PrVD 61.2 35.4-69.8 50.5 10.8 name ‘Guercif barbel’ for this new species.
PrAD 97.7 49.7-107.8 75.9 18.4
PVL 30.4 15.4-34.3 24.3 5.6 HABITAT AND BIOLOGY: The species inhabits
CPL 58.1 30.9-60.6 44.6 9.5 large rivers, mainly in mid-stream, usually in riffle
APL 32.8 19.1-34.8 26.3 4.9 areas. No information exists on the spawning period
BD 26 12.4-28.1 19.5 4.9 or reproductive behaviour.
BLD 14.9 7.8-16.2 11.5 2.7
HL 35 20.5-37.8 27.8 5.6
CONSERVATION: Currently, Luciobarbus guercifen-
sis is a rare species that has been found in few places,
PrOL 17.3 8.1-18.8 12.6 3.2
localized in riffle areas. These riffle areas are becoming
ED 5 3-5.2 4 0.7
scarce in Moulouya River as a consequence of increas-
PsOL 15.4 8.7-16 12.1 2.3 ing crop irrigation at its headwaters. The water taken for
IOW 10.5 5.1-11.3 7.9 1.9 irrigation, as well as the impact of fertilizers and pesti-
B1L 9.7 4.7-11 7.4 2 cides on water quality in the lower courses has likely
B2L 12.8 7.1-13.7 10.2 2.1 been the primary cause of the recent decline of this
PFL 30 16.8-33.3 24.1 4.9 population. No quantification of the decline in num-
VFL 27.3 16.8-31 22.7 4.5 bers is available. We suggest that this species should be
DFL 17.1 9.2-17.7 13.4 2.8 included in the IUCN category of Endangered.
DFH 27.2 15.7-28.5 22 4
GENETICS: Uncorrected-p distance of mitochon-
AFL 11 5.8-12.1 8.6 2.0
drial cytb gene between Luciobarbus guercifensis and
AFH 26.4 16.3-28.5 22.1 3.9
the other analysed species are presented in Table 3.
CFL 32.5 16.5-34.1 26.1 5.3 L. guercifensis shows 63 diagnostic positions in the
LL 45 45-48 45.7 0.9 cytb gene with respect to other rheophilic barbels.
RSA 7.5 5.5-8.5 7.4 0.5
RSB 5.5 5.5 5.5 - Luciobarbus zayanensis sp. nov. Doadrio, Casal-López
& Yahyaoui
http://lsid:zoobank.org:act:7A79DAD4-B60A-43F9-8BF6-
than the postorbital length ( x=12 mm). The dorsal fin
617B34670E64
is posterior on the body but slightly more anterior than
in other rheophilic barbs, with the predorsal distance HOLOTYPE: Fig. 8, Table 6. MNCN 279706, male, 113 mm (SL);
being 46.2-50% of SL. The profile of the dorsal fin Chbouka River, Oum er Rbia Basin, El Herri (Elhri), Atlantic
is concave, with the last single ray ossified with more slope in Morocco (32.859510, -5.621355) (Fig. 1); 27/3/2013.
872 ATSL. Coll. Doadrio, I; Yahyaoui, A; Garzón, P and Perea, S.
than 25 denticulations (Fig. 5). In males, the anal fin
is longer than the one seen in L. nasus and L. magni- PARATYPES: Table 6. MNCN 55094 one specimen from the Douna
atlantis with its height 76-92% of APL. The caudal River, Oum er Rbia Basin, El Khemis, Atlantic slope in Morocco
peduncle is less deep than in L. magniatlantis and (32.750866, -5.541695); 7/5/1988. Coll. Doadrio, I; Merino, M;
Cubo J and González, JL. MNCN 208115-208116: 2 specimens
L. nasus, with a height 9.9-10.9% of SL. The length of
from Serrou River, Oum er Rbia Basin, El Herri, Atlantic slope
the anal caudal peduncle is 2.1 to 2.6 times its height. In in Morocco (32.827621, -5.615255). 27/4/2000. Coll. Doadrio, I;
males, the pectoral and ventral fins are long with pec- Garzón, P; Doadrio, A and Doadrio, I Jr. MNCN 54987-55000: 14
toral fins usually reaching the origin of the ventral fins. specimens from the Arba River, Oum er Rbia Basin, Ksiba, Atlantic
Males exhibited numerous nuptial tubercles of equal slope in Morocco (32.566810, -6.017450). 8/5/1988. Coll. Doadrio, I;
size distributed over the body and fins. The caudal fin Merino, M; Cubo J and González, J.L. MNCN 208168-208169,

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Taxonomy of rheophilic Luciobarbus from Morocco 11

MNCN 71972-71978: 9 specimens from the Oum er Rbia River, El

Borj, Atlantic slope in Morocco Morocco (33.015769, -5.629534).
18/4/1991. Coll. Doadrio, I; Perdices, A and Cubo, J. MNCN
279704-279705, MNCN 279707, MNCN 280083, MNCN 279695-
279696: 6 specimens from the Chbouka River, Oum er Rbia Basin, El
Herri, Atlantic slope in Morocco (32.859510, -5.62135527/3/2013.
27/3/2013. Coll. Doadrio, I; Yahyaoui, A; Garzón, P; and Perea, S.
Holotype and a series of paratypes (33 specimens) have been
Fig. 8.— Holotype of Luciobarbus zayanensis from the Chbouka deposited at the Fish Collection of the Museo Nacional de Ciencias
River, Oum er Rbia Basin. El Herri, Morocco. MNCN 279706. Naturales, Madrid, Spain.
SL=113 mm.
Fig. 8.— Holotipo de Luciobarbus zayanensis del Río Chbouka, DIAGNOSIS: Differs from other known species of
Cuenca del Oum er Rbia. El Herri, Marruecos. MNCN 279706. Luciobarbus by the following combination of char-
SL=113 mm. acters: 49-53 scales along the lateral line ( x =51,
median=51); 8.5-9.5 scales ( x=9.3, median=9.5) above
lateral line and 5.5-7.5 ( x=6.5, median=6.5) below
Table 6.— Morphometric and meristic measurement of lateral line. The last single ray of the dorsal fin is
the holotype and paratypes of Luciobarbus zayanensis. strongly ossified with a maximum width of 12-14.5%
Acronyms are defined in the Material and Methods section. (median=13.9%) of its length. The last single dorsal fin
Tabla 6.— Medidas morfométricas y merísticas del holotipo ray is densely denticulate along its length and, in adults,
y paratipos de Luciobarbus zayanensis. Los acrónimos the number of denticulations is smaller than 23 (Fig. 5).
están definidos en la sección de Material y Métodos. The skull is narrow, with the width of the ethmoid bone
smaller than its length. The inter-orbital distance is
1.2 to 1.9 times (median=1.6) eye diameter. The pha-
Holotype
ryngeal teeth formula in adults is 4.3.2. The number
MNCN 279706 Paratypes n=32
of vertebrae is 41-43 (x=42, n=10). Large deep cau-
Morphometric Measurements Standard
measurement (mm) Range Mean Deviation
dal peduncle 2.1-2.6% of APL. Lachrymal bone with
narrow posterior segment (manubrium). Differences
SL 113 64.5-178.0 118.2 25.1
in diagnostic characters among analysed Luciobarbus
PrDD 56.7 33.3-92.5 59.5 12.7
populations are presented in Table 4.
PrPD 30 17.9-41.5 29.7 5.3
PrVD 54.7 33.3-79.4 55.1 10.5 DESCRIPTION: D III-V 8, A III 5, P I 15-16, V I 6, C
PrAD 78.4 44.6-121.6 81.7 17.2 18; LL 49-53 ( x =51, median=51), RSA 8.5-9.5 ( x =9.3,
PVL 26.4 13.6-42.3 27.1 6.6 median=9.5), RSB 5.5-7.5 ( x =6.5, median=6.5).
CPL 43.7 25.4-76.3 48.5 11.3 Pharyngeal teeth in adults 4.3.2. GR 13-16 ( x =14.5,
APL 28.3 16.0-45.2 29.8 6.3
median=14.5), VE 41-43 ( x =42, n=10). Similarly to
BD 20.8 12.0-43.4 25.7 7.0
other rheophilic Luciobarbus, it is a small-sized spe-
cies, rarely reaching 200 mm SL. Females are larger
BLD 12.3 6.1-20.6 13.3 3.0
( x =128.6 mm) than males ( x =112.9 mm). The body
HL 28.9 17.4-43.4 29.6 5.8 is elongated with maximum body depth 18-24% of
PrOL 14.9 7.8-20.5 13.8 2.8 SL in males and 21-24% in females. The head is typi-
ED 4.6 3.5-6.3 4.7 0.6 cally sharp and narrow. The inter-orbital distance is
PsOL 12 8.8-17.3 12.8 1.9 1.2 to 1.9 times the eye diameter in males and 1.4 to
IOW 7.1 4.1-12.5 7.8 1.8 1.9 times in females. The skull is narrow, with the
B1L 8.1 4.8-8.0 6.5 1.1 ethmoid bone longer than wide. The frontal bones are
B2L 10.5 6.2-11.8 9.4 1.6
narrower relative to other rheophilic barbs, notably in
the anterior part. Infraorbital bones are narrow, and
PFL 23.4 15.1-33.4 24.1 4.1
the lachrymal bone is longer than in other rheophilic
VFL 22.1 14.0-30.6 22.1 3.5
barbs and possesses a long manubrium. The height of
DFL 15.6 8.0-23.0 14.7 3.1 the manubrium is 10-13% the lachrymal bone length.
DFH 25 14.7-32.7 22.8 4.1 The kinethmoid bone is more robust than the one
AFL 8.3 5.3-14.1 9.2 2.1 found in other rheophilic barbs. Thick lips and bar-
AFH 21 11.3-29.2 20.2 4.2 bels with granular appearance. The first pair of bar-
CFL 27.1 15.1-37.5 25.5 4.9 bels is short and positioned at the anterior extreme of
LL 51 49.0-53.0 51.1 1.2 the mouth, reaching the insertion of the second pair
RSA 9.5 8.5-9.5 9.3 0.4
of barbels. The second pair of barbels is thick and
reaches the posterior edge of the eye. The anterior bar-
RSB 5.5 5.5-7.5 6.6 0.6
bel length is 20.1-29.9%, and the second 29.7-39.5%

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12
of HL. The snout is prominent, with preorbital length
10.9-12.1% of SL in females and 10.6-13.2 in males.
The preorbital length ( x =14.8 mm in females and
x =13.5 mm in males) is greater than the postorbital
length ( x =13.5 mm in females and x =12.3 mm in
males). The pectoral and ventral fins are larger in males
than in females. The pectoral fin length is 76-83%
( x =80%) of HL in females and 72-90% ( x =83%)
of HL in males. The ventral fin length is 65.5-74%
( x =70%) of HL in females and 65.5-85% ( x =78%)
of HL in males. The dorsal fin is located posteriorly
on the body, with the predorsal distance being 48.9-
52.5% of SL in females and 48.5-52.5% in males. The
profile of the dorsal fin is concave, with the last single
ray of the dorsal fin ossified with few and strong den-
ticulations (Fig. 5). The caudal peduncle is elongated,
and the least body depth is 11-11.9% SL in females
and 9.5-11.8% SL in males. The length of the anal
caudal peduncle is 2.1 to 2.3 times the height of the
anal peduncle in females and 2.1-2.6 times in males.
Males exhibited numerous equal-sized nuptial tuber-
cles distributed over the body and fins. The caudal fin
length is 20-23% of SL in females and 18.4-24.5%
of SL in males. Morphometric and meristic measure-
ments for the holotype and paratypes of Luciobarbus
­zayanensis are represented in Table 6. The coloration
of L. ­zayanensis is brownish with a paler ventral area
and some black spots along the body (Fig. 2).
DISTRIBUTION: This species is endemic to Oum er
Rbia basin, inhabiting riffle areas with clear waters in
the Atlas Mountains (Fig. 1).
ETYMOLOGY: The species name ‘zayanensis’ has
been selected because it is mainly distributed around
Middle Atlas region, in the area inhabited by the
Zayanes shepherds.
COMMON NAME: We suggest the English common
name ‘Zayan barbel’ for this new species.
HABITAT AND BIOLOGY: The species inhabits
­riffle areas in headwaters of large rivers. Spawning
takes places from late April to early May.
CONSERVATION: The habitat of L. zayanensis is
being transformed by dam and reservoir constructions
and by increased pollution linked to agriculture and
urban development for tourism. The population of
L. zayanensis is in decline, therefore we suggest that
this species should be included in the IUCN category
of Vulnerable.
GENETICS: Uncorrected-p distances of mitochon-
drial gene cytb between L. zayanensis and the other
analysed species are presented in Table 3. Luciobarbus
zayanensis shows 5 diagnostic positions in the cytb
gene.
Graellsia, 72(1), junio 2016, e039 — ISSN-L: 0367-5041 – http://dx.doi.org/10.3989/graellsia.2016.v72.153
I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui
Acknowledgments
Many persons have participated in the field sampling trips.
We warmly thank J. Cubo, M. Merino, J. L. González, P. Garzón,
I. Doadrio Jr., A. Doadrio, A. Perdices, Y. Bernat, and S. El
Gharbi. We would also like to thank L. Alcaraz, for laboratory
work, G. Solís, curator of ichthyological collection, and I. Rey
and B. Álvarez, curators of the DNA collection of the National
Museum of the Natural Sciences (MNCN-CSIC). We also thank
C. Parejo and M. Pérez for her technical assistance in non-
destructive techniques with the computerized tomography scan
at the MNCN-CSIC. James Maclaine, curator of Natural History
Museum, was always willing to help in the study of the L. nasus
type series. We are grateful also to P. Fernández for format edit-
ing, and the Lucidus Consultancy for English editing of this article.
This project was funded by Ministère de l’Enseignement supérieur
from Morocco to the Project NºPPR/2015/2 Impact des change-
ments climatiques sur la diversité génétique des poissons des eaux
douces du Maroc. Permission for fish collection was provided by
Eaux et Foret Commission of Morocco.
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14 I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui

Appendix 1.— Kruskal–Wallis test and Non-parametric Mann-Whitney’s pairwise post hoc
comparisons for all populations. Values of Kruskal-Wallis test (H) below variables. Values of Mann–
Whitney test are below the diagonal. The median is the diagonal of each variable. Significant
differences p<0.05 (*); p<0.01 (**). Acronyms are defined in the Material and Methods section.

Apéndice 1.— Test de Kruskal–Wallis y comparaciones post hoc no paramétricas de Mann-

Whitney para todas la poblaciones. Los valores del test de Kruskal-Wallis (H) aparecen debajo de
las variables. Los valores del test de Mann–Whitney están debajo de la diagonal. La mediana es la
diagonal de cada variable. Diferencias significativas p<0.05 (*); p<0.01 (**). Los acrónimos están
definidos en la sección de Material y Métodos.

Tensift Oum er Rbia Moulouya

Variables Populations (n=14) (n=21) (n=12)
SL (H=0.502) Tensift 108.6
Oum er Rbia 0.449 112.9
Moulouya 0.763 0.968 109.2
PrDD (H=14.63**) Tensift 52.9/0.49
Oum er Rbia 0.88 55.8/0.5
Moulouya <0.0001** <0.0001** 53.3/0.48
PrPD (H=16.54**) Tensift 25.2/0.24
Oum er Rbia 0.005** 29.4/0.26
Moulouya <0.0001** 0.052 28.7/0.26
PrVD (H=4.197) Tensift 50.2/0.46
Oum er Rbia 0.96 54.7/0.47
Moulouya 0.095 0.057 52/0.46
PrAD (H=1.556) Tensift 73.4/0.68
Oum er Rbia 0.827 78.4/0.68
Moulouya 0.529 0.177 77.2/0.7
PVL (H=13.73**) Tensift 24.9/0.24
Oum er Rbia 0.002** 24.9/0.22
Moulouya 0.0005** 0.8427 25/0.22
CPL (H=1.89) Tensift 42.7/0.41
Oum er Rbia 0.372 44.2/0.4
Moulouya 0.978 0.177 44.3/0.41
APL (H=17.8**) Tensift 25.3/0.24
Oum er Rbia <0.0001** 28.3/0.25
Moulouya 0.019 0.06 26/0.24
BD (H=24.63**) Tensift 22.4/0.21
Oum er Rbia 0.259 23.4/0.2
Moulouya <0.0001** <0.0001** 19.7/0.18
BLD (H=34.48**) Tensift 12.8/0.12
Oum er Rbia <0.0001** 12.5/0.11
Moulouya <0.0001** <0.0001** 11.7/0.11
HL (H=29.07**) Tensift 25.1/0.24
Oum er Rbia <0.0001** 28.9/0.25
Moulouya <0.0001** 0.234 28.5/0.25
PrOL (H=2.33) Tensift 12.7/0.12
Oum er Rbia 0.578 13.5/0.12
Moulouya 0.106 0.365 13.6/0.11
PsOL (H=22.8**) Tensift 11.1/0.1
Oum er Rbia <0.0001** 12.3/0.11
Moulouya 0.0002** 8427 12/0.11
ED (H=30.41**) Tensift 4.7/0.04
Oum er Rbia 0.002 4.6/0.04
Moulouya <0.0001** <0.0001** 4.2/0.04

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Taxonomy of rheophilic Luciobarbus from Morocco 15

Appendix 1.— (Continued)

Tensift Oum er Rbia Moulouya

Variables Populations (n=14) (n=21) (n=12)
PFL (H=28.05**) Tensift 20.9/0.2
Oum er Rbia 0.0001** 24.2/0.21
Moulouya <0.0001** 0.0009** 24.5/0.22
VFL (H=24.74**) Tensift 19.4/0.18
Oum er Rbia 0.0009** 22.2/0.2
Moulouya <0.0001** 0.001** 23.5/0.2
DFL (H=3.7) Tensift 13.8/0.12
Oum er Rbia 0.699 14.8/0.13
Moulouya 0.132 0.074 13.6/0.12
DFH (H=26.43**) Tensift 19.6/0.18
Oum er Rbia <0.0001** 23.1/0.2
Moulouya <0.0001** 0.7812 21.9/0.2
AFL (H=1.27) Tensift 8.3/0.08
Oum er Rbia 0.88 9.1/0.08
Moulouya 0.218 0.451 8.9/0.08
AFH (H=25.74**) Tensift 17.6/0.16
Oum er Rbia 0.1623 19.9/0.17
Moulouya <0.0001** <0.0001** 22.4/0.2
IOW (H=32.72**) Tensift 7.9/0.07
Oum er Rbia <0.0001** 7.1/0.06
Moulouya 0.132 <0.0001** 8.1/0.07
LL (H=37.66**) Tensift 49
Oum er Rbia <0.0001** 51
Moulouya <0.0001** <0.0001** 45
RSA (H=40.5**) Tensift 8.5
Oum er Rbia <0.0001** 9.5
Moulouya <0.0001** <0.0001** 7.5
RSB (H=21.65**) Tensift 6.5
Oum er Rbia 0.03* 6.5
Moulouya 0.002 <0.0001** 5.5

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16
Appendix 2.— Osteological features.
Apéndice 2.— Carcaterísticas osteológicas.
Appendix 2-1.— Dorsal view of the skull of the populations
under study. A: Tensift population (N’Fiss River), SL 135.8 mm.
B: Oum er Rbia population (Chbouka river), SL 126.5 mm.
C: Moulouya population (Moulouya River), SL 139 mm.
D: Syntype of Barbus nasus, SL 108.7 mm. Etm: ethmoid bone;
fr: frontal.
Apéndice 2-1.— Vista dorsal del cráneo de las poblaciones
estudiadas. A: población del Tensift (Río N’Fiss), SL 135,8 mm.
B: población del Oum er Rbia (Río Chbouka), SL 126,5 mm.
C: población del Moulouya (Moulouya River), SL 139 mm. D:
Sintipo de Barbus nasus, SL 108,7 mm. Etm: ethmoid; fr: frontal.
Appendix 2-2.— Frontal view of the skull of the populations
under study. A: Tensift population (N’Fiss River), SL 135.8 mm.
B: Oum er Rbia population (Chbouka river) showed a short and
wide kinethmoid bone, SL 126.5 mm. C: Moulouya population
(Moulouya River), SL 139 mm. D: Syntype of Barbus nasus,
SL 108.7 mm. E: Syntype of Barbus nasus, SL 81.5 mm.
Etm: Ethmoid bone; Kim: Kinethmoid bone; Pmx: Premaxilla.
Arrows show the width of kinethmoid bone.
Apéndice 2-2.— Vista frontal del cráneo de las poblaciones
estudiadas. A: población del Tensift (Río N’Fiss), SL 135,8 mm.
B: la población del Oum er Rbia (Río Chbouka) mostró un
hueso kinetmoides corto y ancho, SL 126,5 mm. C: población
del Moulouya (Río Moulouya), SL 139 mm. D: Sintipo de
Barbus. nasus, SL 108,7 mm. E: Sintipo de Barbus nasus, SL
81,5 mm. Etm: Etmoides; Kim: Kinetmoides; Pmx: Premaxilar.
Las flechas muestran la anchura del kinetmoides.
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I. Doadrio, M. Casal-López, S. Perea, & A. Yahyaoui
Appendix 2-3.— Infraorbital bones of the studied populations.
A: Tensift population (N’Fiss River), SL 135.8 mm. B: Oum er
Rbia population (Chbouka river), SL 126.5 mm. C: Moulouya
population (Moulouya River), SL 139 mm. D: Syntype of
Barbus nasus, SL 108.7 mm. E: Syntype of Barbus nasus, SL
81.5 mm. Lcr: Lacrymal. Arrows show the width of lachrymal
manubrium.
Apéndice 2-3.— Huesos infraorbitarios de las poblaciones
estudiadas. A: población del Tensift population (Río N’Fiss),
SL 135,8 mm. B: población del Oum er Rbia (Río Chbouka),
SL 126,5 mm. C: población del Moulouya (Río Moulouya),
SL 139 mm. D: Sintipo de Barbus nasus, SL 108,7 mm.
E: Sintipo de Barbus nasus, SL 81,5 mm. Lcr: Lacrimal. Las
flechas muestran la anchura del manubrio del lacrimal.
Appendix 2-4.— Dentary bone of the studied populations. A:
Tensift population (N’Fiss River), SL 135.8 mm. B: Oum er
Rbia population (Chbouka river), SL 126.5 mm. C: Moulouya
population (Moulouya River), SL 139 mm D: Syntype of Barbus
nasus, SL 108.7 mm. Den: Dentary; An: Anguloarticular.
Apéndice 2-4.— Hueso dentario de las poblaciones
estudiadas. A: población del Tensift (N’Fiss River), SL 135,8
mm. B: población del Oum er Rbia (Río Chbouka), SL 126,5
mm. C: población del Moulouya (Río Moulouya), SL 139 mm
D: Sintipo de Barbus nasus, SL 108,7 mm. Den: Dentario; An:
Anguloarticular.
Taxonomy of rheophilic Luciobarbus from Morocco 17

Appendix 2-7.— External view of the pharyngeal teeth of

Appendix 2-5.— Maxilla of the studied populations. A: Tensift
population (N’Fiss River) showed a short anterior process,
SL 135.8 mm. B: Oum er Rbia population (Chbouka River),
SL 126.5 mm. C: Moulouya population (Moulouya River),
SL 139 mm D: Syntype of Barbus nasus, SL 108.7 mm.
E: Syntype of Barbus nasus, SL 81.5 mm.
Apéndice 2-5.— Maxilar de las poblaciones estudiadas. A: la
población del Tensift (Río N’Fiss) mostró un proceso anterior
corto, SL 135,8 mm. B: población del Oum er Rbia (Río
Chbouka river), SL 126,5 mm. C: población del Moulouya
(Río Moulouya), SL 139 mm D: Sintipo de Barbus nasus,
SL 108,7 mm. E: Sintipo de Barbus nasus, SL 81,5 mm.
the populations studied. Width of the inner branch between
arrows. A: Tensift population (N’Fiss River) showed a flexed
superior branch, SL 135.8 mm. B: Oum er Rbia population
(Chbouka river) showed a strong inner branch, SL 126.5
mm. C: Moulouya population (Moulouya River) showed a thin
pharyngeal bone. SL 139 mm. D: Syntype of Barbus. nasus,
SL 108.7 mm. E: Syntype of Barbus nasus, SL 81.5 mm. ibr:
Inner Branch. obr: Over branch.
Apéndice 2-7.— Vista externa de los dientes faríngeos de
las poblaciones estudiadas. Anchura de la rama interna entre
flechas. A: población del Tensift (Río N’Fiss) mostró una rama
superior flexionada, SL 135,8 mm. B: población del Oum er
Rbia (Río Chbouka) mostró una rama interna robusta, SL
126,5 mm. C: población del Moulouya (Río Moulouya) mostró
un hueso faríngeo delgado, SL 139 mm. D: Sintipo de Barbus
nasus, SL 108,7 mm. E: Sintipo de Barbus nasus, SL 81,5
mm. ibr: Rama interna. obr: Rama superior.

Appendix 2-6.— Pharyngeal teeth of the populations studied.

A: Tensift population (N’Fiss River) showed 5 teeth in the
external row, 3 in the middle, and 2 in the inner row, SL 135.8.
B: Oum er Rbia population (Chbouka river) showed 4 teeth
in the external row, 3 in the middle, and 2 in the inner row, SL
126.5 mm. C: Moulouya population (Moulouya River) showed
4 teeth in the external row, 2 in the middle, and one in the inner
row, SL 139 mm. D: Syntype of Barbus nasus showed 4.3.2
pharyngeal teeth, SL 81.5 mm.
Apéndice 2-6.— Dientes faríngeos de las poblaciones
estudiadas. A: población del Tensift (Río N’Fiss) mostró 5
dientes en la fila externa, 3 en la del medio y 2 en la fila interna,
SL 135,8 mm. B: población del Oum er Rbia (Río Chbouka)
mostró 4 dientes en la fila externa, 3 en la del medio y 2 en
la fila interna, SL 126,5 mm. C: población del Moulouya (Río
Moulouya) mostró 4 dientes en la fila externa, 2 en la del medio
y una en la fila interna, SL 139 mm. D: Sintipo de Barbus nasus
mostró 4.3.2 dientes faríngeos, SL 81,5 mm.

Graellsia, 72(1), junio 2016, e039 — ISSN-L: 0367-5041 – http://dx.doi.org/10.3989/graellsia.2016.v72.153

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